Scaling-up the biodiversity–ecosystem functioning relationship: the effect of environmental heterogeneity on transgressive overyielding

Abstract

Knowledge of how biodiversity sustains ecosystem function comes predominantly from studies focused on small spatial scales. Thus, we know relatively little about the role of biodiversity at larger scales of space and time where habitats become increasingly heterogeneous. Efforts to upscale the relationship between biodiversity and function have yielded inconclusive results. Given that increasing habitat heterogeneity is a ubiquitous consequence of increasing spatial scale, we asked: as habitat heterogeneity increases, can single species continue to maintain ecosystem function? Or, does transgressive overyielding (functioning of species mixture divided by the functioning of the highest functioning single species) change with habitat heterogeneity? We addressed this using a combination of computer simulations, an experiment and a meta-analysis. The three parts followed the same rationale: habitat heterogeneity was increased by aggregating local habitats with different conditions into larger and more heterogeneous landscapes. The computer simulations showed that, on average, transgressive overyielding increased with habitat heterogeneity because monoculture functioning decreased with habitat heterogeneity. We tested this expectation experimentally by varying the strain richness from one to five species across 10800 bacterial communities in five different habitats defined by sub-inhibitory concentrations of antibiotics. On average, the experimental results concurred with the simulations. We tested the generality of this result using a meta-analysis of 26 published experiments that manipulated habitat conditions and species richness. This confirmed that transgressive overyielding tended to increase with habitat heterogeneity but only when species were specialised to different habitats and were not inhibited in mixtures by negative species interactions. This was not the case in several experiments used in our meta-analysis where one species maximised functioning across all habitats, contrary to the assumptions of many ecological models. Our results illustrate the importance of biodiversity at larger spatial scales with more heterogeneity but also highlights contingencies that this pattern depends on.